Kent Kainulainen

Deep divergences in the coffee family and the systematic position of Acranthera

Despite extensive efforts, there are unresolved questions on evolutionary relationships in the angiosperm family Rubiaceae. Here, information from six loci and 149 Rubiaceae taxa provide new insights. Acranthera and Coptosapelta are strongly supported as sisters. Pollen grains of Acranthera possess several features common in Rubiaceae, but amongst potential similarities with the unusual grains of Coptosapelta are the nature of the apertures and the structure of the sexine. Luculia, Acranthera and Coptosapelta are excluded from the three subfamilies Ixoroideae, Cinchonoideae and Rubioideae. Sipaneeae and Condamineeae form a clade, sister to remaining Ixoroideae. Rondeletieae and Guettardeae are sisters to remaining Cinchonoideae. Colletoecema is sister to remaining Rubioideae, followed by the Urophylleae–Ophiorrhizeae clade. Nuclear ITS provided structured information at all phylogenetic levels, but the main gain from adding nrITS was the increased resolution. Average support values also increased but were generally high also without nrITS and the increase was not statistically significant.

A Revised Time Tree of the Asterids : Establishing a Temporal Framework For Evolutionary Studies of the Coffee Family (Rubiaceae)

Divergence time analyses in the coffee family (Rubiaceae) have all relied on the same Gentianales crown group age estimate, reported by an earlier analysis of the asterids, for defining the upper age bound of the root node in their analyses. However, not only did the asterid analysis suffer from several analytical shortcomings, but the estimate itself has been used in highly inconsistent ways in these Rubiaceae analyses. Based on the original data, we here reanalyze the divergence times of the asterids using relaxed-clock models and 14 fossil-based minimum age constraints. We also expand the data set to include an additional 67 taxa from Rubiaceae sampled across all three subfamilies recognized in the family. Three analyses are conducted: a separate analysis of the asterids, which completely mirrors the original asterid analysis in terms of taxon sample and data; a separate analysis of the Gentianales, where the result from the first analysis is used for defining a secondary root calibration point; and a combined analysis where all taxa are analyzed simultaneously. Results are presented in the form of a time-calibrated phylogeny, and age estimates for asterid groups, Gentianales, and major groups of Rubiaceae are compared and discussed in relation to previously published estimates. Our updated age estimates for major groups of Rubiaceae provide a significant step forward towards the long term goal of establishing a robust temporal framework for the divergence of this biologically diverse and fascinating group of plants.

Phylogenetic structure and clade circumscriptions in the Gardenieae complex (Rubiaceae)

In this study we investigate the large and diverse Rubiaceae-Gardenieae and closely related tribes Bertiereae, Coffeeae, Cremasporeae, Octotropideae, and Pavetteae. Some of the tribes or groups have been shown to be monophyletic and strongly supported, but the phylogeny of this large complex is still far from being satisfactorily elucidated particularly for Gardenieae, both in terms of intertribal relationships as well as tribal delimitations. We reconstruct the phylogeny of the complex using an extensive sampling of 108 genera and five plastid DNA regions. Phylogenetic relationships demonstrate that Gardenieae sensu Andreasen & Bremer is polyphyletic, as Burchellia, Didymosalpinx, Monosalpinx, and Mantalania are closer to Octotropideae-Cremasporeae. In addition, Pavetteae and the investigated members of Aulacocalyceae are nested in a supported but partially unresolved Gardenieae-Pavetteae clade. Within this clade, several strongly supported groups are resolved: an Aidia group, an Alibertia group, a Gardenia group, Pavetteae including Pelagodendron, a Porterandia group, a Randia group, a Rothmannia group (including Aulacocalyx and Heinsenia), a Sherbournia group, and the two isolated genera Massularia and Schumanniophyton. The latter genus presented a high rate of genetic substitutions, which resulted in perturbations of the phylogenetic reconstruction. A revised tribal circumscription is given for Gardenieae, the Alibertia group is recognized at tribal level as an emended Cordiereae, and a new tribe, Sherbournieae, is described to accommodate the members of the Sherbournia group.

Croton aleuritoides P.E. Berry (Euphorbiaceae), a Distinctive New Tree Species from Montagne des Français in Northern Madagascar

Abstract Berry, P.E., B. Van Ee, K. Kainulainen & H. Razafindraibe (2016). Croton aleuritoides P.E. Berry (Euphorbiaceae), a distinctive new tree species from Montagne des Français in northern Madagascar. Candollea 71: 181–188. In English, English and French abstracts. Croton aleuritoides P.E. Berry (Euphorbiaceae) is described from deciduous forests on tsingy limestone in the Antso River basin of Montagne des Français in northern Madagascar. This remarkable new species is distinguished by its arborescent habit; large, broadly ovate-cordate, deciduous leaves; adaxial epipetiolar glands; sparsely lepidote pubescence; elongate stipules; large seeds with a whitish fleshy covering when fresh; and cinerescent fruits that are indehiscent when dispersing from the tree and only dehiscing much later on the ground. It is remarkable that such a large tree from a relatively small massif close to the regional capital has gone undetected for so long. With its very limited area of occupancy and small population size, it should be considered an endangered species.

Phylogeny of Euclinia and allied genera of Gardenieae (Rubiaceae), and description of Melanoxerus, an endemic genus of Madagascar

We performed molecular phylogenetic analyses of the Randia clade of the tribe Gardenieae using both plastid and nuclear DNA data. In the phylogenetic hypotheses presented, the African genera Calochone, Euclinia, Macrosphyra, Oligocodon, Pleiocoryne, and Preussiodora are resolved as a monophyletic group. Support is also found for a clade of the Neotropical genera Casasia, Randia, Rosenbergiodendron, Sphinctanthus, and Tocoyena. This Neotropical clade is resolved as sister group to the African clade in analyses of combined plastid and nuclear data. The genus Euclinia appears polyphyletic. The species described from Madagascar represent an independent lineage, the position of which is moreover found to be incongruent between datasets. Plastid and ribosomal DNA data support a sister-group relationship to the mainland African clade, whereas the low-copy nuclear gene Xdh supports a closer relationship to the Neotropical genera. The phylogenetic reconstructions also indicate that Casasia and Randia are not monophyletic as presently circumscribed. A taxonomic proposal is made for the recognition of the Malagasy taxon at generic level as Melanoxerus.

Croton cupreolepis (Euphorbiaceae), a New Coppery-Lepidote Tree Species from Eastern Madagascar

Abstract Croton cupreolepis (Euphorbiaceae) is newly described from montane, evergreen forests of eastern Madagascar. It is distinguished by its coppery-lepidote trichomes that form a dense covering on the floral buds and inflorescences, as well as on the dorsal side of the petals of both pistillate and staminate flowers, and also by its distinctive recurved pedicels. The young leaves are covered by the same type of coppery trichomes on the lower surface, but as the leaves expand, the coppery scales become scattered on a background of silvery scales. The species has been confused with Croton chrysodaphne, C. nobilis, or else specimens have been left determined just to genus. Its broad latitudinal distribution in Madagascar suggests that it may once have been more widespread, but has been a victim of the widespread deforestation of primary forest along the eastern slopes of the island.

New species and species reports of Croton L. (Euphorbiaceae) from the eastern forest corridor of Madagascar

Abstract Kainulainen, K., B. Van EE, P. Antilahimena, H. Razafindraibe & P.E.Berry (2016). New species and species reports of Croton L. (Euphorbiaceae) from the eastern forest corridor of Madagascar. Candollea 71: 327–356. In English, English and French abstracts. Six species of Croton from the Moramanga District of the Alaotra-Mangoro Region of eastern Madagascar (Toamasina Prov.) are newly described here, five of which occur in the Ambatovy mining concession. A seventh new species is described from the Ankerana Forest in the Atsinanana Region, some 80 km to the northeast of the town of Moramanga, which is one of the offset areas intended to mitigate the deforestation incurred by the Ambatovy mining project. Of the new species, Croton ferricretus Kainul., B.W. van Ee & P.E. Berry is the one most closely associated with the ultramafic soils where nickel and cobalt is being extracted. Although it is locally common, it is only known from the mine concession. Croton enigmaticus P.E. Berry & B.W. van Ee is considerably less common on the mine concession but is also known from two other sites. Croton droguetioides Kainul. & Radcl.-Sm. is known from Ambatovy and three other areas in the Alaotra-Mangoro Region, and Croton radiatus P.E. Berry & Kainul. is known only from forests on the Ambatovy mining concession and from the area of Zahamena National Park. Croton indrisilvae Kainul., B.W. van Ee & P.E. Berry is a small-leaved species known only from Analamazaotra National Park, and Croton ankeranae Kainul. is another small-leaved species known so far only from Ankerana. Croton plurispicatus P.E. Berry, Kainul. & B.W. van Ee is a distinctive small tree known mainly from the Lakato and Vohibe forests in the southern part of the Ankeniheny-Zahamena Corridor and one other location east of Analamazaotra. Croton hypochalibaeus Baill. is reinstated from synonymy and is now known from the Ambatovy area as well as in several forest remnants within the “tampoketsa” vegetation on the high central plateau of Madagascar, and in canyons of Isalo National Park. An additional species (Croton lasiopyrus Baill.) is identified from the Moramanga area for the first time, but is not currently known from Ambatovy. Amended descriptions are provided for both Croton hypochalibaeus and Croton lasiopyrus, and lectotypes are designated.

A Nomenclator of Croton (Euphorbiaceae) in Madagascar, the Comoros Archipelago, and the Mascarene Islands

Abstract All published names of Croton from Madagascar, the Comoros, and the Mascarenes are treated here. We indicate which names are currently accepted (123 native species and 1 introduced), which ones we consider to be heterotypic synonyms (188), which ones are doubtful (25), and which ones should be excluded (5). We newly designate lectotypes for 108 names, and epitypes for C. anisatus Baill., C. nobilis Baill., and C. submetallicus Baill. A total of 133 names are newly treated as synonyms. One new combination is made, Croton basaltorum (Leandri) P.E.Berry for C. antanosiensis var. basaltorum Leandri, and one new name is proposed, Croton toliarensis B.W.vanEe & Kainul. for C. tranomarensis var. rosmarinifolius Radcl.-Sm.

Rediscovery of two species of Croton (Euphorbiaceae) from littoral habitats of eastern Madagascar

Abstract Kainulainen, K., P.E. Berry & B. Van Ee (2017). Rediscovery of two species of Croton (Euphorbiaceae) from littoral habitats of eastern Madagascar. Candollea 72: 35–44. In English, English and French abstracts. Two species of Croton L. (Euphorbiaceae) previously reported only from their type specimens, Croton chapelieri Baill. and Croton vatomandrensis Leandri, are reevaluated here and found to be geographically widespread along the eastern coastline of Madagascar. We provide updated descriptions, lists of selected additional specimens, distribution maps, and photographs of both species. We also designate an epitype for Croton chapelieri and a lectotype for Croton vatomandrensis. Croton chapelieri was the source of considerable taxonomic confusion, and we treat as synonyms here for the first time Croton aymoninorum Leandri, Croton louvelii Leandri, Croton daphniphylloides Radcl.-Sm., Croton daphniphyllum Radcl.-Sm., Croton daphniphyllum var. hirsutus Radcl.-Sm., and Croton rhododendroides Radcl-Sm. Croton chapelieri var. longepetiolata Radcl-Sm. is considered to be a synonym of Croton submetallicus Baill. With our improved understanding of the geographical ranges of Croton chapelieri and Croton vatomandrensis, we provide IUCN Red List conservation assessments for both species.

A revision of the Adenophorus Group and other glandular-leaved species of Croton (Euphorbiaceae) from northern Madagascar and Mayotte, including three new species

Abstract Kainulainen, K., B. Van Ee, H. Razafindraibe & P.E. Berry (2017). A revision of the Adenophorus Group and other glandular-leaved species of Croton (Euphorbiaceae) from northern Madagascar and Mayotte, including three new species. Candollea 72: 371–402. In English, English abstract. DOI: http://dx.doi.org/10.15553/c2017v722a15 This paper provides a revision of the species of Croton L. (Euphorbiaceae) from northern Madagascar (mainly Antsiranana and northern Mahajanga Provinces) and Mayotte in the Comoros Archipelago that bear glands on the undersides of the leaves. This includes members of the Adenophorus Group as well as two species that do not belong to that group, namely Croton nudatus Baill. and Croton stanneus Baill. Croton nudatus has been a problematic name since its publication due to the leafless state of its type. We were able to match the type with material recently collected near the type locality, and we provide an amended description for this species. Croton stanneus is another previously poorly understood species, which is actually one of the most widespread species of Croton in Madagascar. We also re-evaluate species delimitations in members of the Adenophorus Group from northern Madagascar and Mayotte, and we describe three new species, Croton mayottae P.E. Berry & Kainul., Croton orangeae Kainul. & P.E. Berry, and Croton sahafariensis Kainul. & P.E. Berry. Croton loucoubensis Baill. is resurrected from synonymy, and nine other species names and eight varieties are reduced to synonymy under Croton adenophorus Baill., Croton loucoubensis, Croton nudatus, Croton scoriarum Leandri, Croton stanneus, or Croton tsiampiensis Leandri. Full descriptions are provided for these species as well as for Croton bathianus Leandri, another misunderstood species in the group. A key to the seven species of the Adenophorus Group and the two look-alikes from northern Madagascar and Mayotte is provided. An epitype is designated for Croton nudatus, and lectotypes are designated for Croton adenophorus, Croton bathianus, Croton loucoubensis, Croton scoriarum, Croton stanneus, and Croton tsiampiensis. Received: July 10, 2017; Accepted: August 23, 2017; First published online: September 27, 2017